Age at maturity and diversification in woody angiosperms

Angiosperm diversification has been associated with plant-animal interactions such as seed dispersal and pollination and life-history characters such as rapid growth and fast reproduction. This paper relates a life-history character (age at maturity) to woody angiosperm diversification. Here I present a comparative analysis of data drawn from the literature, indicating that time to first reproduction is shorter in woody angiosperms than in gymnosperms. In addition, age at maturity is negatively correlated with the rate of diversification (measured as the number of species per genus) at all the taxonomic levels analyzed and also when phylogenetically independent contrasts were conducted. This correlation suggests that early reproduction promotes diversification in woody angiosperms. Furthermore, this correlation is not a confounding effect of the association between age at maturity and other ecological factors that promote angiosperm diversification, such as pollination and seed dispersal systems.     

Community-level flowering phenology and fruit set: Comparative study of 25 woody species in a secondary forest in Japan

We investigated the flowering phenology, pollinator visitation, and fruit set of 25 animal-pollinated woody species in a warm temperate secondary forest in Japan. Various species flowered sequentially from February to October. The principal pollinators were bumblebees, honey-bees, flies and/or beetles and birds; bumblebees and flies/beetles pollinated most trees. The duration of flowering was shorter for species that bloomed in the middle of the season than it was for species that bloomed earlier or later in the season. The timing of flowering was more synchronous within species that had a shorter flowering duration; this was also detected when phylogenetically independent contrasts were calculated. This could be important for the effective pollination of species with a short flowering duration because such species bloom sequentially over a short period of less than 1month around May. Fruit set was related not to pollinator type, sex expression, flowering sequence (in order of the date of peak flowering) or flowering duration, but to the relative abundance of the species in the forest. This correlation was detected for fly- and beetle-pollinated species but not for bumblebee-pollinated species. Thus, relatively rare plant species with opportunistic pollinators might experience limited fruit set because of insufficient pollinator services. Bagging experiments conducted on eight hermaphrodite species revealed that the fruit set of bagged flowers was nearly zero, lower than that of control flowers. These results indicate the importance of pollinators for successful reproduction and thus for the coexistence of plants in this secondary forest.     

传粉方式和果实类型对木本植物繁殖物候的影响

该研究利用谱系独立比较法(Phylogenetically Independent Contrasts,PIC)和Wilcoxon秩和检验法,分析中国科学院植物研究所植物园(39°54'N,116°12'E)中的84个物种170株个体的传粉方式和果实类型对木本开花时间和结实时间的影响,其中Wilcoxon秩和检验法检验的结果作为PIC检验结果的参考。结果表明:(1)传粉方式显著影响植物开花和结实时间,风媒花植物比虫媒花植物开花和结实早;(2)果实类型对结实时间的影响在考虑和不考虑物种间系统发育关系时表现不同,当不考虑物种间系统发育关系时,肉质果实植物结实时间比非肉质果实植物早;(3)不同的传粉方式间以及不同的果实类型间植物的花果间隙期无显著差异,但本研究结果显示肉质果实植物结实时间比非肉质果实植物大约早20d。由此推论:(1)植物固有属性,如传粉方式和果实类型,会影响植物繁殖物候,且不同的属性影响强度不同;(2)与某一特定物候期或繁殖器官相关性大的属性对该物候期的影响可能更大,如传粉方式对开花时间的影响可能大于其对结实时间的影响,而果实类型对结实物候期的影响更大。     

Phenotypic selection on flowering phenology and size in two dioecious plant species with different pollen vectors

Dioecious plants may be pollinated biotically by animals or abiotically via wind or water currents. It has been hypothesized that these two types of pollen vectors might impose contrasting selective pressures on plant flowering phenology. In the present study we describe the flowering phenology of two sympatric dioecious species with contrasting pollination modes: Mercurialis perennis (wind‐pollinated) and Tamus communis (insect‐pollinated). We estimated selection differentials and gradients for flowering time and flowering synchrony. As flowering time might depend on the accumulation of enough internal resources, we also estimated direct and indirect selection on plant size. Both species have male‐biased sexual ratios, and males are bigger and produce larger flower displays than females, but only in T. communis do males bloom earlier and for longer than females. Selection gradients suggest that selection tends to favor early‐flowering females of T. communis. There is no evidence of direct current selection on the flowering phenology of M. perennis. Intersexual differences in phenology fit with sex allocation and sexual selection theories. As we hypothesized, phenology of the animal‐pollinated species is under stronger selection than that of the wind‐pollinated species and we discuss the potential role of pollen vectors in shaping the flowering phenologies of the study species.     

石斛属植物叶片生物量与叶片养分含量间的关联进化

叶片生物量和元素含量是显著影响叶片生理功能的两类重要的叶片经济性状, 且二者紧密相关。然而过去的研究多数关注C、N和P三种元素含量, 而对其他元素含量关注较少。维管附生植物不能直接从土壤中吸收养分而经常遭受养分胁迫, 养分含量与生物量之间的协同关系在这类植物中表现得尤为突出。选择维管附生石斛属植物为研究对象, 应用系统发育独立对比分析, 检测了叶片生物量和元素含量的系统发育保守性以及这些性状间的关联进化。结果表明, 叶片干物质含量、比叶重、C含量、N含量、P含量以及C/N具有较强的系统发育信号, 并且叶片干物质含量和比叶重与这四个元素含量性状间存在较强的关联进化。另外, 经系统发育校准后, Ca含量和Mg含量间存在显著正相关; P含量与Si含量间存在显著正相关; Zn含量与N含量、N/P以及Mn含量间存在显著正相关, 而与C/N间存在显著负相关。结果为探讨维管附生植物叶片生物量和元素含量的主要影响因素, 以及从进化角度探讨二者在植物进化适应过程中的生态协同功能提供案例, 对进一步了解维管附生植物的功能适应性具有重要意义。     

西安和宝鸡木本植物花期物候变化及温度敏感度对比

植物物候是指示生态系统对气候变化响应的重要证据。已有研究多基于代表性站点的物候观测数据研究物候特征及其对气候变化的响应规律。同一气候区内,不同站点的物候变化及对温度变化响应的敏感度是否一致仍需深入探讨。本文选择同属于暖温带湿润区汾渭平原气候区的西安和宝鸡为研究区,利用"中国物候观测网"在两个站点21个共有物种的开花始期和开花末期数据,比较了1987—2016年两站点各植物花期物候变化特征及其对温度变化响应的敏感度差异。结果表明,西安和宝鸡各物种的开花始期和开花末期均以提前趋势为主。大部分物种开花始期在西安的提前趋势(平均趋势-0.57 d/a)明显强于在宝鸡的提前趋势(平均趋势-0.29 d/a),但开花末期趋势差异不显著。除紫薇和迎春的敏感度差异较大外,其他物种开花始期和开花末期的温度敏感度在两站点间非常接近,无显著差异。由此可见,在同一气候区的不同站点,因增温幅度不同,植物的始花期变化存在较大差异,不能用单站点的物候变化反映整个气候区的物候变化。但同一植物在单站点的温度敏感度可以较好的反映同一气候区其他站点的植物物候-气候关系。本文研究结果可为利用有限站点的物候观测数据分析区域物候...     

Differences in seedling growth behaviour among species: trait correlations across species, and trait shifts along nutrient compared to rainfall gradients

1 Species-pairs from woody dicot lineages were chosen as phylogenetically independent contrasts (PICs) to represent evolutionary divergences along gradients of rainfall and nutrient stress, and within particular habitat types, in New South Wales, Australia. Seedlings were grown under controlled, favourable conditions and measurements were made for various growth, morphological and allocation traits.
2 Trait correlations across all species were identified, particularly with respect to seedling relative growth rate (RGR) and specific leaf area (SLA), a fundamental measure of allocation strategy that reflects the light-capture area deployed per unit of photosynthate invested in leaves.
3 Across all species, SLA, specific root length (SRL) and seed reserve mass were the strongest predictors of seedling RGR. That is, a syndrome of leaf and root surface maximization and low seed mass was typical of high RGR plants. This may be a high-risk strategy for individual seedlings, but one presumably mitigated by a larger number of seedlings being produced, increasing the chance that at least one will find itself in a favourable situation.
4 Syndromes of repeated attribute divergence were identified in the two sets of gradient PICs. Species from lower resource habitats generally had lower SLA. Thus, in this important respect the two gradients appeared to be variants of a more general 'stress' gradient.
5 However, trends in biomass allocation, tissue density, root morphology and seed reserve mass differed between gradients. While SLA and RGR tended to shift together along gradients and in within-habitat PICs, no single attribute emerged as the common, primary factor driving RGR divergences within contrasts. Within-habitat attribute shifts were of similar magnitude to those along gradients.     

Effects of experimental shifts in flowering phenology on plant-pollinator interactions

Climate change has led to phenological shifts in flowering plants and insect pollinators, causing concern that these shifts will disrupt plant-pollinator mutualisms. We experimentally investigated how shifts in flowering onset affect pollinator visitation for 14 native perennial plant species, six of which have exhibited shifts to earlier flowering over the last 70 years and eight of which have not. We manipulated flowering onset in greenhouses and then observed pollinator visitation in the field. Five of six species with historically advanced flowering received more visits when flowering was experimentally advanced, whereas seven of eight species with historically unchanged flowering received fewer visits when flowering earlier. This pattern suggests that species unconstrained by pollinators have advanced their flowering, whereas species constrained by pollinators have not. In contrast to current concern about phenological mismatches disrupting plant-pollinator mutualisms, mismatches at the onset of flowering are not occurring for most of our study species.     

Plant hydraulics and photosynthesis of 34 woody species from different successional stages of subtropical forests

It is important to understand the ecophysiological characters of plants when exploring mechanisms underlying species substitution in the process of plant succession. In the present study, we selected 34 woody species from different stages of secondary succession in subtropical forests of southern China, and measured their hydraulic conductivity, gas exchange rates, leaf nutrients and drought‐tolerance traits such as xylem resistance to cavitation, turgor loss point and carbon isotope ratio. Principal component analysis revealed that early‐, mid‐ and late‐successional species were significantly separated along axis 1, which was strongly associated with hydraulic‐photosynthetic coordination. In contrast to species distributed in late‐successional forest, early‐successional species had the highest hydraulic conductivity, net photosynthetic rates, photosynthetic nitrogen and phosphorus use efficiencies, but had the lowest photosynthetic water‐use efficiency. However, changes of the measured drought‐tolerance traits of the 34 species along the succession did not demonstrate a clear trend – no significant correlations between these traits and plant successional stages were found. Moreover, the trade‐off between hydraulic efficiency and safety was not identified. Taken together, our results suggested that hydraulic efficiency and photosynthetic function, rather than drought tolerance, play an important role in species distributions along plant succession in subtropical forests.     

Density, body mass and parasite species richness of terrestrial mammals

We investigated the relationships between helminth species richness and body mass and density of terrestrial mammals. Cross-species analysis and the phylogenetically independent contrast method produced different results. A non-phylogenetic approach (cross-species comparisons) led to the conclusion that parasite richness is linked to host body size. However, an analysis using phylogenetically independent contrasts showed no relationship between host body size and parasite richness. Conversely, a non-phylogenetic approach generated a negative relationship between parasite richness and host density, whereas the independent contrast method showed the opposite trend – that is, parasite richness is positively correlated with host density. From an evolutionary perspective, our results suggest that opportunities for parasite colonization depend more closely on how many hosts are available in a given area than on how large the hosts are. From an epidemiological point of view, our results confirm theoretical models which assume that host density is linked to the opportunity of a parasite to invade a population of hosts. Our findings also suggest that parasitism may be a cost associated with host density. Finally, we provide some support for the non-linear allometry between density and mammal body mass (Silva and Downing, 1995), and explain why host density and host body mass do not relate equally to parasite species richness.     

Testing the link between the latitudinal gradient in species richness and rates of molecular evolution

Numerous hypotheses have been proposed to explain latitudinal gradients in species richness, but all are subject to ongoing debate. Here we examine Rohde's (1978, 1992) hypothesis, which proposes that climatic conditions at low latitudes lead to elevated rates of speciation. This hypothesis predicts that rates of molecular evolution should increase towards lower latitudes, but this prediction has never been tested. We discuss potential links between rates of molecular evolution and latitudinal diversity gradients, and present the first test of latitudinal variation in rates of molecular evolution. Using 45 phylogenetically independent, latitudinally separated pairs of bird species and higher taxa, we compare rates of evolution of two mitochondrial genes and DNA-DNA hybridization distances. We find no support for an effect of latitude on rate of molecular evolution. This result casts doubt on the generality of a key component of Rohde's hypothesis linking climate and speciation.     

Plant reproductive phenology over four years including an episode of general flowering in a lowland dipterocarp forest,Sarawak, Malaysia

The first systematic observation of a general flowering, a phenomenon unique to lowland mixed-dipterocarp forests in Southeast Asia, is presented. During general flowering, which occurs at irregular intervals of 3–10 yr, nearly all dipterocarp species together with species of other families come heavily into flower. We monitored reproductive phenology of 576 individual plants representing 305 species in 56 families in Sarawak, Malaysia. Observations continued for 53 mo from August 1992 and covered one episode of a general flowering cycle. Among 527 effective reproductive events during 43 mo, 57% were concentrated in the general flowering period (GFP) of 10 mo in 1996. We classified 257 species into flowering types based on timing and frequency of flowering. The most abundant type was “general flowering” (35%), which flowered only during GFP. The others were “supra-annual” (19%), “annual” (13%), and “sub-annual” (5%) types. General flowering type and temporal aggregation in reproductive events were commonly found among species in various categories of taxonomic groups, life forms, pollination systems, and fruit types. Possible causes for general flowering, such as promotion of pollination brought about by interspecific synchronization and paucity of climatic cues suitable for flowering trigger, are proposed, in addition to the predator satiation hypothesis of Janzen (1974) .     

开花时间与伴生种对鹤首马先蒿传粉和生殖成功的影响

开花物候是物种间相互作用的重要生活史特征和适合度因子,在全球气候变化的背景下而备受关注.为探讨开花时间如何存种内和种间水平上影响植物的传粉和生殖成功,我们连续3年(2003-2005)对不同花期和伴生种存在情况下的鹤首马先蒿(Pedicularis gruina)的传粉者访花忠实度、受粉率、坐果率、单果种子产量和果实被啃食频率进行了比较研究.结果表明鹤首马先蒿的坐果率主要受其传粉环境的影响:在没有伴生种时,不同时期鹤首马先蒿的坐果率没有显著差异,34-38％的花可以坐果;在有伴生种存在时既可以显著提高其坐果率,也可显著降低其坐果率,这取决于传粉者类型以及伴生种密穗马先蒿(P.densispica)花期的差异.密穗马先蒿具有花蜜和花粉双重报酬,在群落中可以作为主体物种吸引传粉者并间接促进与其伴生的鹤首马先蒿的传粉和生殖成功.同样无蜜的管花马先蒿(P.siphonantha)和鹤首马先蒿伴生,则是通过提高群落水平对传粉者的吸引力进而直接促进鹤首马先蒿的传粉和生殖成功.此外,研究结果也表明开花时间对坐果率没有显著影响,但是显著影响单果种子产量和果实被啃食的频率.在相同的传粉条件下早花期植株单果种子产量显著高于晚花期的种子产量,同时早花期的果实被啃食的频率显著增加.     

Prediction of flowering time in Brassica oleracea using a quantitative trait loci-based phenology model

Uniformly developing plants with a predictable time to harvest or flowering under unfavourable climate conditions are a major breeding goal in crop species. The main flowering regulators and their response to environmental signals have been identified in Arabidopsis thaliana and homologues of flowering genes have been mapped in many crop species. However, it remains unclear which genes determine within and across genotype flowering time variability in Brassica oleracea and how genetic flowering time regulation is influenced by environmental factors. The goal of this study is model-based prediction of flowering time in a B. oleracea DH-line population using genotype-specific and quantitative trait loci (QTL) model input parameters. A QTL-based phenology model accounting for genotypic differences in temperature responses during vernalisation and non-temperature-sensitive durations from floral transition to flowering was evaluated in two field trials. The model was parameterised using original genotype-specific model input parameters and QTL effects. The genotype-specific model parameterisation showed accurate predictability of flowering time if floral induction was promoted by low temperature (R(2) = 0.81); unfavourably high temperatures reduced predictability (R(2) = 0.65). Replacing original model input parameters by QTL effects reduced the capability of the model to describe across-genotype variability (R(2) = 0.59 and 0.50). Flowering time was highly correlated with a model parameter accounting for vernalisation effects. Within-genotype variability was significantly correlated with the same parameter if temperature during the inductive phase was high. We conclude that flowering time variability across genotypes was largely due to differences in vernalisation response, although it has been shown elsewhere that the candidate FLOWERING LOCUS C (FLC) did not co-segregate with flowering time in the same population. FLC independent vernalisation pathways have been described for several species, but not yet for B. oleracea.     

Adaptation to climate through flowering phenology: a case study in Medicago truncatula

Local climatic conditions likely constitute an important selective pressure on genes underlying important fitness‐related traits such as flowering time, and in many species, flowering phenology and climatic gradients strongly covary. To test whether climate shapes the genetic variation on flowering time genes and to identify candidate flowering genes involved in the adaptation to environmental heterogeneity, we used a large Medicago truncatula core collection to examine the association between nucleotide polymorphisms at 224 candidate genes and both climate variables and flowering phenotypes. Unlike genome‐wide studies, candidate gene approaches are expected to enrich for the number of meaningful trait associations because they specifically target genes that are known to affect the trait of interest. We found that flowering time mediates adaptation to climatic conditions mainly by variation at genes located upstream in the flowering pathways, close to the environmental stimuli. Variables related to the annual precipitation regime reflected selective constraints on flowering time genes better than the other variables tested (temperature, altitude, latitude or longitude). By comparing phenotype and climate associations, we identified 12 flowering genes as the most promising candidates responsible for phenological adaptation to climate. Four of these genes were located in the known flowering time QTL region on chromosome 7. However, climate and flowering associations also highlighted largely distinct gene sets, suggesting different genetic architectures for adaptation to climate and flowering onset.